A thorny issue: Woody plant defence and growth in an East African savanna

Recent work suggests that savanna woody plant species utilise two different strategies based on their defences against herbivory; a low nutrient/high chemical defence strategy and a nutrition paired with mostly architectural defences strategy. The concept that chemical and structural defences can augment each other and do not necessarily trade-off has emanated from this work. In this study, we examine woody plant defence strategies, how these respond to herbivore removal and how they affect plant growth in an East African savanna. At three paired long-term exclosure sites with high browser and mixed-feeder densities at Mpala Ranch, Kenya, we investigated: (a) whether defences employed by the dominant fine- and broad-leaved woody savanna species form defence strategies and if these align with previously proposed strategies, (b) how nine key plant defence traits respond to herbivore removal and (c) how effective the different defence strategies are at protecting against intense herbivory (by measuring plant growth with and without herbivores present). We identified three defence strategies. We found a group (a) with high N, short spines and high N-free secondary metabolites, a group (b) with high N, long spines and low N-free secondary metabolites and a group (c) with moderate N, no spines and low N-free secondary metabolites (most likely defended by unmeasured chemical defences). Structural defences (spine length, branching) were generally found to be induced by herbivory, leaf available N increased or did not respond, and N-free secondary metabolites decreased or did not respond to herbivory. Species with long spines combined with increased “caginess” (dense canopy architecture arising from complex arrangement of numerous woody and spiny axis categories) of branches, maintained the highest growth under intense browsing, compared to species with short spines and high N-free secondary metabolites and species with no spines and low N-free secondary metabolites. Synthesis. At our study site, structural traits (i.e. spines, increased caginess) were the most inducible and effective defences against intense mammalian herbivory. We propose that high levels of variability in the way that nutrient and defence traits combine may contribute to the coexistence of closely related species comprising savanna woody communities

Differential effects of defoliation by mopane caterpillars and pruning by African elephants on the regrowth of Colophospermum mopane foliage

Plant responses to herbivory vary depending on herbivory type, yet the comparative effects of defoliation (e.g. by insects) and pruning (e.g. by large mammals) on a single tree species are poorly documented. We investigated this in the Northern Province of South Africa by comparing the regrowth of Colophospermum mopane trees previously defoliated by caterpillars or pruned by elephants, the two main browsers of C. mopane foliage. Shoots were up to 160% and 125% longer after natural (elephant) and simulated pruning and leaves ~25% longer in regrowth after natural pruning (n = 13–15 trees per treatment). Shoot density and chemical defences in leaves (tannin:protein ratio and total polyphenolic concentration) were, however, no different from control trees. Simulated defoliation resulted in statistically insignificant changes to regrowth in terms of leaf and shoot size (both slightly decreased) and shoot density (slightly increased). Natural (caterpillar) defoliation, however, resulted in regrowth with significantly decreased shoot and leaf size (about 50% and 20% of control lengths, respectively), as well as decreased leaf chemical defence. Shoot and leaf length were longer on trees flushing for the first time after pruning and late-season defoliation had a greater negative impact than mid-season defoliation. Despite the differences in regrowth characteristics after pruning and defoliation, mopane plants showed no apparent trade-off in investment between tolerance and resistance after either herbivory type, as neither regrowth nor chemical defence occurred at the expense of the other

Intraspecific host preferences of mopane moths (Imbrasia belina) in mopane (Colophospermum mopane) woodland

Colophospermum mopane trees are a common larval food plant of the mopane moth (Imbrasia belina), which commonly reaches outbreak proportions. Our study investigated factors determining host tree choice by ovipositing mopane moths within a single host species (C. mopane). Tree size was the primary determinant of oviposition at both the habitat (tall riverine-, medium height woodland- and short-shrub mopane) and individual tree scale, with an increase in the number of egg masses with increasing tree size (estimated by height and canopy volume). Preference for larger trees was only evident at the habitat scale, however, as individual trees were utilized as expected according to the availability of their canopy size class. More detailed tree characteristics, such as leaf size, shoot size, stem number and even leaf chemistry (protein : tannin ratio and total polyphenols) had no influence on host choice. Host choice was based on the most obvious measure of resource abundance, namely tree size, as expected for an outbreak species, since resource availability rather than nutritional quality is likely to be the primary determinant of larval survival

Seasonal effects of Rhus lancea and Celtis africana on intake preference and physiological responses in South African indigenous goats

This study investigated the seasonal effects of Rhus lancea and Celtis africana leaves on preference, intake, weight, and serum metabolites in South African indigenous, mature, male goats. Twelve mature, male goats weighing 34 ± 5.9 kg (mean ± SD) were randomly allocated to two groups of six and kept in metabolic crates for periods of 21 days in October 2015 and March, May, and August of 2016. A browser diet of R. lancea and C. africana and a control diet (lucerne and concentrates) were randomly allocated to each group. Measurements taken included nutritional composition of browse per season, and browse preference, intake, weight changes, and serum metabolites in the goats. The acid detergent fibre (24–36%) and neutral detergent fibre (26.9–70.4 %) in R. lancea over the months were greater than in C. africana (50.3–53.2% and 49.4–55.4%, respectively). In the preference study, the goats preferred C. africana more in October (51.2 vs 48.8%), March (51.4 vs 48.6%), and May (54.3 vs 45.7%). Goats on the browser diet lost weight in March, May, and in August whereas those on the control diet gained weight. The serum urea concentration of goats consuming browser diets in May and August (1.8–3.3 mmol/l) was lower than the normal range, consistent with animals failing to derive their protein requirements from the diet. Goats prefer to browse C. africana more than R. lancea. The study also indicated the need for supplementation to meet maintenance requirements in animals fed R. lancea and C. africana.College of Agriculture and Environmental Science

Native legume species as potential fodder crops in the mediterranean renosterveld shrubland, South Africa

The limited grazeable size of renosterveld vegetation in the Western Cape warranted assessment of chemical and fibre composition of the species to determine their nutritive value. We hypothesised that Fabaceae has chemically superior palatable species relative to Poaceae, Asteraceae and Asparagaceae species in the area. Plant samples were collected in the Overberg region from 11 sites. The species sampled from these four families Fabaceae, Poaceae, Asteraceae, and Asparagaceae were analysed for macro- and micro-elements, fibre and secondary compounds. At each site, replicated soil samples were also collected and analysed for pH, macro- and micro-elements. Fabaceae recorded the greatest concentration of crude protein, but the Poaceae was the most fibrous relative to Fabaceae and Asteraceae. All the species contained adequate levels of nutrients except P, Zn and sometimes Cu. Soil concentration of macronutrients and carbon in the soil varied with location. Based on crude protein, neutral detergent fibre and digestibility, the three most important parameters defining forage quality, the Fabaceae species Aspalathus hispida, A. angustifolia, A. nigra and A. submissa were regarded as superior forage species in the Mediterranean renosterveld. Livestock relying on only the renosterveld species in the Western Cape as forages should receive P and Zn supplements

Stem growth of woody species at the Nkuhlu exclosures, Kruger National Park: 2006–2010

An important aspect of managing African conservation areas involves understanding how large herbivores affect woody plant growth. Yet, data on growth rates of woody species in savannas are scarce, despite its critical importance for developing models to guide ecosystem management. What effect do browsing and season have on woody stem growth? Assuming no growth happens in the dry season, browsing should reduce stem growth in the wet season only. Secondly, do functional species groups differ in stem growth? For example, assuming fine-leaved, spiny species’ growth is not compromised by carbon-based chemical defences, they should grow faster than broad-leaved, chemically defended species. Dendrometers were fixed at 20 cm in height on the main stems of 244 random plants of six woody species in three plots (all large herbivores excluded, partial exclusion, and control) and observed from late 2006 to early 2010. Average monthly increment (AMI) per dendrometer and season (dry, wet) was calculated and the interaction between plot and season tested per species, controlling for initial stem girth. AMIs of Combretum apiculatum, Dichrostachys cinerea and Grewia flavescens were zero in the dry season, whilst those of Acacia exuvialis, Acacia grandicornuta and Euclea divinorum were either positive or negative in the dry season. Wet-season AMI of D. cinerea and dry-season AMI of G. flavescens tended to be reduced by browser exclusion. Net AMI (sum of the seasonal AMIs) was tested per species, but results suggested that only D. cinerea tended to be affected by browser exclusion. The results also suggested that stem radial growth of some fast-growing species is more prone to reduction by browser exclusion than the growth of other species, potentially reducing their competitiveness and increasing their risk of extirpation. Finally, the usefulness of grouping woody species into simple functional groups (e.g. fine-leaved vs. broad-leaved) for ecosystem management purposes in savannas requires further consideration. Conservation implications: Growth rates of woody plants are important parameters in savanna models, but data are scarce. Monitoring dendrometers in manipulative situations over several years can help fill that gap. Results of such studies can be used to identify species prone to high risk of extirpation